WHAT IS EVOLUTION?

Evolution is the process by which all living things have developed from primitive organisms through changes occurring over billions of years, a progression that includes the most advanced animals and plants. Biologists agree that all living things arose through a long history of changes shaped by physical and chemical processes that are still taking place.

PHYLOGENIC TREE OF PLANT EVOLUTION Evidence suggests that plants evolved from green algae Plants - multicellular, photosynthetic eukaryotes that range in size from the duckweed to the giant coastal redwoods Important ecologically, industrially, and medically

Believed to have evolved from freshwater green algae over 500 million years ago Evidence - Both green algae and plants Contain chlorophylls a and b and various accessory pigments Store excess carbohydrates as starch Have cellulose in their cell wall

The evolution of plants is marked by four innovations Nourish and protect a multicellular embryo that completes the life cycle Have vascular tissue transporting water and solute to cells when the plant body is surrounded by air Produce seeds that contain an embryo and stored organic nutrients within a protective coat Flowers that attract pollinators, such as insects, and give rise to fruits, food for animals to help disperse the seeds

I. Overview of land plant evolution A. Four main groups of land plants Bryophytes Pteridophytes Gymnosperms and Angiosperms

Groups are distinguished from algae by reproduction (life cycle) that involves the development of a multi-cellular embryo attached to the mother plant for its protection and nourishment.

1. Bryophytes liverworts, hornworts, mosses - Bryophytes have no vascular tissues. The rest three groups are all vascular plants. Vascular plants have cells that are joined to produce tubes that transport water and nutrients throughout the plant.

Bryophytes live in damp/moist environments and are small so they dont need vascular tissue. They are sometimes called non-vascular plants. Algae that we saw in last chapter live in water and dont need vascular tissue because nutrients come from surrounding water.

The vascular plants are, in order of their evolution: 2. Pteridophytes ferns, horsetails, lycophytes a. seedless plants 3. Gymnosperms conifers, ginkgo, cycads, gnetopsids a. early seed plants b. produce naked seeds 4. Angiosperms flowering plants

a. seeds protected by growing in ovaries b. majority of modern plants are in this group

B. Charophyceans - green algae most closely related to land plants.

1. Both charophyceans and land plants are multi-cellular, eukaryotic, photoautotrophs.

2. Both have rosette cellulose-synthesizing complexes rose-shaped arrays of proteins that synthesize the cellulose components that make up plant cell walls. Because all these features are shared between the groups, both land plants and charophyceans must have a common ancestor. Close algal relatives of plants (Charophyceans: Chara (top), Coleochaete orbicularis (bottom)

In order to grow on land, the land plants needed to evolve terrestrial adaptations to survive. C. Terrestrial adaptations can be used to distinguish land plants from charophycean algae. These adaptations are: 1. Apical meristems

2. Multi-cellular, dependent embryos 3. Alternation of generations 4. Spore walls contain sporopollenin 5. Multi-cellular gametangia

1. Apical meristems a. localized areas of cell division at tips of roots and shoots. Apical meristems of plant shoots and roots

2. Multi-cellular, dependent embryos

a. Embryo develops within female tissue; female plant provides nutrition (sugars, proteins).

b. Placental transfer cells that enhance the transfer of nutrients from the parent to the embryo. 3. Alternation of generations Two multi-cellular body forms: A sporophyte represents the diploid generation and a gametophyte represents the haploid generation Sporophyte (2n) produces spores by meiosis A spore is a haploid reproductive cell that develops into a new organism without needing to fuse with another reproductive cell A spore undergoes mitosis to become a gametophyte

Gametophyte (n) produces gametes In plants, eggs and sperm are produced by mitosis A sperm and egg fuse, forming a diploid zygote that undergoes mitosis and becomes the sporophyte

4. Spore walls contain sporopollenin

a. Sporopollenin is a polymer that makes the walls of plant spores very tough and resistant to harsh conditions.

b. Sporopollenin is the most durable organic material known.

c. Spores are produced by sporangia (cells in the sporophyte) through the process of meiosis.

d. Durable spores are an adaptation for surviving on land. Can withstand long periods of adverse conditions.

Easily transported by wind and water.

A fern spore Reproductive Adaptation to the Land Environment Sporophyte dominance can be associated with an increasing adaptation for reproduction in a dry, terrestrial environment Ferns are seedless vascular plants with a dominant sporophyte The sporophyte produces spores that disperse (scatter) separate gametophytes The gametophyte is a small, heart-shaped structure that has no vascular tissue and can dry out if the environment is not moist Each archegonium on the surface of a gametophyte produces an egg that is fertilized by a flagellated sperm, which must swim to the archegonium in a film of external water

6. Other terrestrial adaptations common to many land plants a. Epidermis covered by a waxy cuticle to prevent excess loss of water. Pores (stomata) in cell layer can be opened and closed to allow O2 out and CO2 in. Cuticle - relatively impermeable layer and provides an effective barrier to water loss, but it also limits gas exchange Leaves have little openings called stomata (sing., stoma) that let carbon dioxide enter while allowing oxygen and water to exit

b. Except for bryophytes, land plants have vascular tissue in roots, stems, and leaves.

- Xylem consists of dead cells that carry water and nutrients from roots to the rest of the plant.

- Phloem consists of living cells that distribute sugars and amino acids throughout the plant.

II. Origin of land plants

A. Theory is that land plants evolved from charophycean algae over 500 million years ago.

Evidence: 1. Homologous chloroplasts

2. Homologous cell walls made of cellulose

3. Homologous peroxisomes

4. Similar DNA sequences

B. Alternation of generations in plants may have originated by delayed meiosis

Zygote Sporophyte Many, many spores

1. Occurs on land because its more difficult to produce zygotes. (No water for swimming sperm)

2. By producing sporophyte, many gametophytes can be produced from one zygote because many, many spores are produced. This maximizes output of sexual reproduction.

C. Adaptations to shallow water pre-adapted plants for living on land

1. Charophycean algae inhabit shallow waters and need to survive when water levels drop. Lead to increasing ability to survive entirely on dry land.

III. Bryophytes

A. Gametophyte is the dominant generation in the life cycles of bryophytes

B. Life cycle of bryophytes

1. Bryophyte sporophytes produce and disperse huge numbers of spores.

C. Ecological and economic benefits of bryophytes

1. Bryophytes were the worlds only plants for 100 million years.

2. Peat bogs are made mostly of moss called sphagnum. They contain 400 billion tons of carbon and cut down the amount of greenhouse gases. Peat is harvested, dried, and used as a fuel.

3. Sphagnum is harvested for use as a soil conditioner and plant packing material. IV. Origin of vascular plants - Pteridophytes = ferns - Gymnosperms = fir trees - Angiosperms = flowering plants

A. Vascular plants evolved additional terrestrial adaptations

1. Xylem and phloem 2. Dominant sporophyte generation independent of the gametophyte Different from the bryophytes

B. Cooksonia evolved over 400 million years ago oldest known vascular plant

Cooksonia, a vascular plant of the Silurian

V. Pteridophytes: seedless vascular plants A. Pteridophytes provide clues to evolution of roots and leaves

1. There is evidence that roots evolved from subterranean portions of stems.

2. There are two types of leaves:

a. Leaves of lycophytes are microphylls. Microphylls are small leaves with a single, unbranched vein.

b. Leaves of other modern vascular plants are megaphylls. Megaphylls are typically larger and have a branched vascular system.

Hypotheses for the evolution of leaves

B. Sporophyte-dominant life cycle evolved in seedless vascular plants (Pteridophytes)

1. Alternation of generations

2. Dominant sporophyte versus dominant gametophyte in bryophytes.

3. Plants are dispersed to new environments as spores; no seeds present The life cycle of a fern

C. Importance of Pteridophytes

1. Dominant plants in Carboniferous period

2. Extensive beds of coal from these plants

Carboniferous forest based on fossil evidence

Overview of seed plant evolution There were three (3) major reproductive adaptations in the evolution of seed plants. 1) Reduction of gametophyte (Gametophytes of seed plants are microscopic, so they can exist within the sporophyte). 2) Seeds became an important means of dispersal. 3) Pollen (Sperm cells) eliminated the liquid-water requirement for fertilization.

1) Reduction of the gametophyte continued with the evolution of seed plants. In seed plants, the very small female (1N) gametophyte develops from spores retained in the sporangia of the (2N) sporophyte. Why?? Delicate female gametophytes are protected from environmental stress. Embryos that are produced are also protected from stress.

In flowering plants: o Sporophyte produces seeds that disperse separate sporophytes protected by seed coats Female gametophyte is microscopic and retained and protected within an ovule, a structure located within the tissue of a flower The male gametophytes are pollen grains that are transported by wind, insects, or birds They do not need external water to reach the egg

A Reduction in the size of the gametophyte as sporophyte becomes dominant

Three variations on gametophyte/sporophyte relationships

2) Seeds became an important means of dispersal.

Seeds are resistant to environmental stress, and are multi-cellular complex structures that consist of a sporophyte embryo, a food supply, and a protective coat.

The embryo develops from the fertilized egg (derived from a spore) retained within the sporangium. The sperm (derived from a spore) is carried (produced by) a pollen grain released from a sporangium. All seed plants are heterosporous. Female spores give rise to female gametes; male spores give rise to male gametes.

The megasporangia produce megaspores (female) and the microsporangia produce microspores (male).

The megasporangium is enveloped by layers of tissue called integuments. The whole structure (integuments + megasporangium) is called the ovule.

3) Pollen (with sperm cells) eliminated the liquid-water requirement for fertilization.

Pollen travels by air or on animals. It eliminates the need for water to be present during fertilization.

Gymnosperms Gymnosperms are vascular plants that bear naked seeds seeds not enclosed in specialized chambers. Mesozoic era was the age of gymnosperms Gymnosperms were the most common plants during the Mesozoic era (the age of dinosaurs).

Four phyla of extant gymnosperms

Phylum Ginkgophyta: e.g., Ginkgo biloba. Phylum Cycadophyta: e.g., cycads. Phylum Gnetophyta: e.g., ephedra

 Phylum Coniferophyta: e.g., pines, firs, spruces

Life cycle of a pine demonstrates the key reproductive adaptations of seed plants

1. The tree is the sporophyte.

2. Female gametophyte develops within the sporangium.

3. Pollen cone has microsporangium that develops into pollen (male gametophyte).

4. After fertilization, the embryo develops and is surrounded by food reserves and a seed coat.

5. Embryo grows to produce a new sporophyte.

Angiosperms are flowering plants that form seeds inside a protective chamber called an ovary. Examples: Monocots: Orchids, lilies, grasses, palms, bamboo (Eu)dicots: Daisies, maples, snapdragon, pea, oaks Angiosperms (Flowering Plants) Systematists are identifying angiosperm clades. Originally, only:

Monocots are angiosperms that possess one embryonic seed leaf (cotyledon).

Dicots are angiosperms that possess two embryonic seed leaves (cotyledons).

A comparison of monocots and dicots

The flower is the defining reproductive adaptation of angiosperms

Flowers are made up of four types of modified leaves sepals, petals, stamens, and carpels.

a. Stamens are the male reproductive organs that produce microspores.

b. Carpels are female reproductive organs that produce megaspores.

The structure of a flower.

A fruit is a mature ovary

Protect dormant seeds.

Fruit aids in seed dispersal. - Wind dispersal - Attachment and transportation - Consumption berries contain seeds to be feces Fruit adaptations that enhance seed dispersal passed in

Life cycle of an angiosperm

Mature plant is the sporophyte

Some plants easily self-pollinate, but most have mechanisms to ensure crosspollination.

Male gametophytes reach female gametophytes by producing a pollen tube.

Next slide: The life cycle of an angiosperm.

Angiosperms dominated the earth at the end of the Mesozoic era

Radiation of angiosperms represents the transition from Mesozoic to Cenozoic

Angiosperms and animals have affected one anothers evolution

Coevolution is the mutual influence on the evolution of two different species interacting with each other and reciprocally influencing each others adaptations.

e.g., Pollinator-plant relationships